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And a Rhinocerotoid Perissodactyl from the Middle Eocene Santiago Formation of Southern California

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And a Rhinocerotoid Perissodactyl from the Middle Eocene Santiago Formation of Southern California Paludicola 11(1):8-30 October 2016 © by the Rochester Institute of Vertebrate Paleontology NEW RECORDS OF EPIHIPPUS (EQUIDAE) AND A RHINOCEROTOID PERISSODACTYL FROM THE MIDDLE EOCENE SANTIAGO FORMATION OF SOUTHERN CALIFORNIA Thomas S. Kelly1 and Paul C. Murphey2 1Research Associate, Vertebrate Paleontology Department, Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles, California 90007 2Research Associate, Department of Paleontology, San Diego Museum of Natural History, 1788 El Prado, San Diego, California 92101 ABSTRACT Records of the middle Eocene equid Epihippus from the West Coast of North America are extremely rare. Previously, only one specimen was reported from California in a faunal list of the Mission del Oro Local Fauna from the Santiago Formation of San Diego County, but this specimen was never described or illustrated and its specific status remained undetermined. Subsequently, additional specimens of Epihippus from southern California have been discovered as the result of paleontological mitigation programs conducted for commercial development projects. Here we provide detailed accounts of all of the Epihippus specimens currently known from California, wherein we recognize the occurrence of two species, E. gracilis and E. uintensis. A rhinocerotoid, which likely represents either Uintaceras or Teletaceras, is also recorded from the Santiago Formation of San Diego County. In addition, a new middle Eocene fauna is documented, the Rancho Mission Viejo Local Fauna, which is the first fauna of the early Uintan North American Land Mammal Age recorded from Orange County. INTRODUCTION Occurrences of middle Eocene horses are extremely rare in southern California. Walsh (1996b) Contrary to the early and late Eocene faunas of reported the first record of Epihippus in a faunal list North America, horses are a relatively rare of the Mission del Oro Local Fauna from the component in middle Eocene faunas of the Santiago Formation of San Diego County, but did not Bridgerian, Uintan and Duchesnean North American provide a description or illustration of the specimen. Land Mammal Ages, primarily being known from the Extensive, ongoing paleontological mitigation western interior and Gulf Coast Plain (Granger, 1908; programs have been conducted since 1996 in Matthew, 1929; McGrew, 1953; Gazin, 1976; southern California, which have resulted in new MacFadden, 1980,1998; Rasmussen, et al., 1999a, records of Epihippus from three additional localities 1999b; Froehlich, 2002; Kelly et al., 2012). During in the Santiago Formation (Figure 1). A the early Eocene Wasatchian North American Land rhinocerotoid specimen, which was not described or Mammal Age, horses were common and diverse, illustrated, was previously recorded from the including at least six genera and 13 species Santiago Formation of San Diego County and (Froehlich, 2002). By the beginning of the misidentified as Hyracodon sp. (Tomiya, 2012). Bridgerian, horse diversity had rapidly declined with These perissodactyl records are highly significant to only one genus, Orohippus Marsh, 1872, being our understanding of the middle Eocene recognized (MacFadden, 1998). Orohippus gave rise biostratigraphy of southern California. The purpose to Epihippus in the earliest Uintan and the latter gave of this report is to provide detailed accounts of all of rise to Duchesnehippus Peterson, 1931, in the the known specimens of Epihippus from California Duchesnean. During the late Duchesnean and reevaluate the taxonomic status of the Duchesnehippus gave rise to Mesohippus Marsh, rhinocerotoid specimen. 1875, which became abundantly represented by numerous species during the early Chadronian (late METHODS Eocene) through earliest Arikareean (early late Oligocene) North American Land Mammal Ages Measurements of teeth were made with an (MacFadden, 1998). optical micrometer to the nearest 0.01 mm. Equid dental terminology follows Froehlich (2002) and that 8 KELLY AND MURPHEY—EPIHIPPUS FROM SOUTHERN CALIFORNIA 9 time scale); N, number of specimens; R, right; SD, standard deviation; tr, greatest transverse width; tra, anterior transverse width; trp, posterior transverse width. Abbreviations for institutions and specimens cited in text are: OCPC, Orange County Paleontological Collection at the Dr. John D. Cooper Archaeological and Paleontological Center, Orange County, California; SDNHM, San Diego Natural History Museum; SDSNH, San Diego Society of Natural History. GEOLOGIC SETTING The type Santiago Formation was first named by Woodring and Popenoe (1945) for outcrops on Santiago Creek to the east of Irvine Park in the Santa Ana Mountains of Orange County, California. The Santiago Formation unconformably overlies the Paleocene Silverado Formation and unconformably underlies the middle Eocene (late Uintan) to Miocene Sespe Formation and/or the informal unit referred to by Whistler and Lander (2003) as the Sespe- Vaqueros Formations undifferentiated (see also Woodring and Popenoe, 1945; Morton, 1974; Schoellhamer et al., 1981; Prothero, 2001). In the type area, the formation is divided into two informal units, the lower and upper beds (Schoelhammer et al., 1981). The lower beds are characterized by medium FIGURE 1. Map of southern California coastal area showing to coarse grained, cross-bedded sandstones, greenish- geographic locations of middle Eocene localities that yielded Epihippus. 1, Rancho Mission Viejo (locality OCPC 03976), gray to reddish mudstones and conglomerate lenses Orange County. 2, Mission del Oro (locality SDSNH 3574), San near the base, whereas the upper beds are Diego County. 3, Ocean Ranch (locality SDSNH 4730), San characterized by medium to fine grained, poorly Diego County. 4, Legoland (locality SDSNH 4131), San Diego bedded sandstones, greenish-gray to reddish County. Insert map shows area covered by map. ________________________________________________ variegated sandy siltstones and claystones (Woodring and Popenoe, 1945; Morton, 1974). for Rhinocerotoidea follows Dashzeveg and Hooker In northern San Diego County, outcrops have (1997) except that we prefer the term cristid obliqua also been referred to the Santiago Formation by instead of metalophid for the cristid that extends Wilson (1972), wherein he recognized three informal anterolingually from the hypoconid to protolophid. members (members A, B and C). Based upon early Upper and lower teeth are designated by uppercase Uintan mammal faunas from the upper part of and lowercase letters, respectively. Open member B and correlation with Calcareous nomenclature qualifiers follow Bengtson (1988). All Nannoplankton Zones, Planktonic Foraminiferal specimens described here are curated in the research Zones, Benthic Foraminiferal Stages and West Coast collections at the Department of Paleontology at the Molluscan "Stages," Walsh (1996b) regarded San Diego Museum of Natural History, San Diego, member B of the Santiago Formation in northwestern California and the Orange County Paleontological San Diego County to span the late Bridgerian to early Collection at the Dr. John D. Cooper Archaeological Uintan North American Land Mammal Ages, or and Paleontological Center, Orange County, about 48-45.5 Ma, and a correlative to the La Jolla California. Detailed locality data are available at Group of southwestern San Diego County. Member these institutions. C of the Santiago Formation is separated from Subzones or biochrons of the Bridgerian and member B by an unconformity in northwestern San Uintan North American Land Mammal Ages (e.g., Diego County and has yielded middle Eocene Br3, Ui1a, Ui1b, Ui2, and Ui3) follow Gunnell et al. mammal faunas of late Uintan and early Duchesnean (2009). Ages (Walsh, 1996b; Walsh et al., 1996; Prothero, Abbreviations are: ap, greatest anteroposterior 2001; Mihlbachler and Deméré, 2009). Based on length; CV, coefficient of variation; L, left; Ma, magnetostratigraphy, Walsh et al. (1996) correlated megannum (one million years in the radioisotopic the La Jolla Group to Chron C21n and the lower third 10 PALUDICOLA, VOL. 11, NO. 1, 2016 of Chron C20r of the Global Polarity Time Scale, or SDSNH 43478, we reexamined it and provide a about 47.9-45.5 Ma. Prothero (2001) reported that detailed description below, which indicates that it the lower part of the type Santiago Formation in cannot be assigned to either Triplopus or Hyracodon. Orange County was entirely within a paleomagnetic _________________________________________ normal zone, which he correlated to Chron C21n. Prothero (2001) was unable to sample the magnetostratigraphy of the upper part of the type Santiago Formation, so its polarity has not been determined. The Bridgerian-Uintan transition is regarded by most recent investigators as occurring within Chron C21n at about 47 Ma (e.g., Walsh, et al., 1996; Robinson et al., 2004; Gunnell, et al., 2009; Kelly and Murphey, 2016). LOCALITIES AND FAUNAS Mission del Oro—Walsh (1996b) named the Mission del Oro Local Fauna based on a fauna from three localities (SDSNH 3570, 3572, 3574) in member C of the Santiago Formation that were discovered during the grading of the Mission del Oro housing project, Oceanside, California (Figure 2, Table 1). The localities occur in brownish, gritty mudstones interbedded within coarse-grained, arkosic channel sandstones, about 9 to 10 m above the disconformity that separates members B and C. Locality SDSNH 3574 yielded the first record of Epihippus from California. Based primarily on the presence of Protylopus sp., cf. P. robustus Golz, 1976, in the
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